The Small Scavenger Guild of Massachusetts.
Continuing forensic taphonomic research has indicated the important role of scavengers in the consumption of soft tissue, bone destruction, and bone dispersal for human remains deposited in terrestrial surface environments (Haglund et al. 1988; Haynes 1983; Pokines 2014, 2016). These scavenging species often include birds and small mammals in the size range of raccoons (Procyon lotor), red fox (Vulpes vulpes), bobcats (Lynx rufus), fisher (Martes pennanti), and rodents, and the role of these small species in the scavenging of large vertebrate remains in terrestrial environments is an under-researched aspect of forensic taphonomy (Jeong et al. 2016; Klippel & Synstelien 2007; Pokines & Baker 2014; Reeves 2009; Rippley et al. 2012; Spradley et al. 2012; Young et al. 2014, 2015, 2016). While larger scavengers such as black bear (Ursus americanus) can be excluded from more urbanized areas and small patches of wild habitat, smaller scavengers can still inhabit these areas and utilize human remains as a food resource. Their feeding may cause bone damage much in the same manner that large species such as coyotes (Canis latrans), wolves (C. lupus), and larger breeds of domesticated dog (C. familiaris) can (Murmann et al. 2006; Pobiner 2008; Pokines 2014), although the gnawed areas may be more limited in scope and/or take longer to manifest. Actualistic studies are needed to address these questions on a regional basis, as the local species represented and population densities will vary greatly.
The ability of small scavengers to disperse remains away from the initial point of deposition is also largely unknown, likely due to the inherent difficulties in observing and tracking such activities over weeks or months. Bone dispersal may play as large a role in bone loss from a skeleton as actual destruction of elements does (Pokines 2015a; Spradley et al. 2012; Young et al. 2016). The increasing distance of isolated bones or fragments from a main concentration of remains makes their recovery far more difficult, as the area to be searched increases as a square function of the distance of dispersal. Komar and Beattie (1998) noted that corvids (crows and their relatives) can disperse small bones over 600 m during normal scavenging activity, and New World vultures (Cathartidae) also have been detected dispersing large vertebrate bone (Reeves 2009; Spradley et al. 2012; Stolen 2003). It is likely that small mammals are also important dispersers of bone, given the propensity of large mammals to transport bone away from the point of initial deposition during feeding activities (Pokines 2014).
It is important to note that the effects of scavenging on a single set of vertebrate remains deposited into a terrestrial surface environment are usually the cumulative effects of multiple species. In ecological terms, a "guild" is a group of species that exploit the same environmental resource in similar ways (Cortes-Avizanda et al. 2012; O'Brien et al. 2010; Olson et al. 2016; Root 1967; Simberloff & Dayan 1991; Wallace & Temple 1987). Small vertebrate species, including mammals and birds, that scavenge decomposing carcasses fit this category, provided that they are utilizing similar portions of the carcass. In the present research, none of the species involved can crush long bones to access the marrow directly and are largely restricted to the soft tissue still attached, although the guild includes some carnivoran and rodent scavengers that can gnaw through areas of thin cortical bone to access interior soft tissues. Many of these species are also exploiting the remains of species larger than they could hunt, so their ecological interactions extend beyond predation to taxa in the size class of humans. These species also would be expected to scavenge a human body introduced into this environment, so analysis of the structure and foraging habits of this guild is necessary for understanding the taphonomic history of forensic scenes.
Previous research in different Nearctic environments has highlighted the potential importance of the small scavenging guild in its effects upon outdoor forensic scenes with human remains. This includes research at the Anthropological Research Facility (ARF) at the University of Tennessee, Knoxville. The placement of fences and their general maintenance prevented most large scavengers of the area from entering the compound where human bodies were continuously placed for decomposition. The fence could not fully exclude smaller scavengers, especially raccoon, which were a nightly visitor to scavenge the human remains (Synstelien 2015). Other small scavengers that frequented the ARF included Virginia opossum (Didelphis virginiana), brown rat (Rattus norvegicus), and white-footed mouse (Peromyscus leucopus). Virginia opossum tended to feed upon the maggots colonizing the human bodies, brown rats were attracted to soft tissue and grease contained in bones, white-footed mice were attracted to soft tissue, and eastern gray squirrels were attracted to dry bone for gnawing (Synstelien 2015). Multiple other potential scavengers were noted, but these were rarely observed directly scavenging the human remains; these taxa include domesticated dog, red fox, domesticated cat (Felis catus), and a variety of bird species.
Komar and Beattie (1998) found that Corvidae were important scavengers on pig (Sus scrofa) carcasses in an environment in Alberta, Canada, and the rodents deer mouse (Peromyscus maniculatus) and southern red-backed vole (Clethrionomys glareolus) as well as Passerine (songbirds, etc.) species were also noted in association with experimental carcasses. Morton and Lord (2006) noted the importance of multiple small scavengers upon pig carcasses in Virginia. Moss (2012) monitored twelve cadavers in a Texas environment over the course of 18 months using game cameras and found multiple small scavengers feeding on the remains. These included American crow (Corvus brachyrhynchos), turkey vulture (Cathartes aura), black vulture (Coragyps atratus), raccoon, Virginia opossum, bobcat, red-shouldered hawk (Buteo lineatus), and gray fox (Urocyon cinereoargenteus). Research in the Palearctic (Young et al. 2014, 2015, 2016) and Australasia (O'Brien et al. 2010) also has indicated the potential importance of the small scavenger guilds in those zoogeographic zones.
The present research was performed at the Outdoor Research Facility (ORF) located in Holliston, Massachusetts, and maintained by the Forensic Anthropology Program, Boston University School of Medicine. The ORF is 32 acres of suburban woodlot bisected by a series of artificial former cranberry bogs that now are more natural wetlands habitat. The tree cover is mixed pine and deciduous species, with small areas of open canopy. Common tree species identified at the ORF include eastern white pine (Pinus strobus), sugar maple (Acer saccharum), black maple (Acer nigrum), American mountain ash (Sorbus americana), American beech (Fagus grandifolia), white oak (Quercus alba), and paper birch (Betula papyrifera). Much of the surrounding region is similarly forested (DeGraaf et al. 2006). Adjacent properties include suburban homes, a cemetery to the east, and a secondary school across the street to the west (Figure 1). The overall facility, however, does not have fenced boundaries, and wildlife is free to roam through the area. The facility has been in use for multiple years and includes two research structures, some fenced-in areas, and an enclosed decomposition field for taphonomic research (Junod 2013; Ricketts 2013; Smith 2014). Sporadic human disturbance occurs within this area, but it is frequented by multiple wildlife species attracted to the plant cover and water resources away from major human disturbances. The surrounding region has multiple small towns but no major urban areas, so many species of wildlife can freely disperse through this area. White-tailed deer (Odocoileus virginianus) are frequent residents, as are coyotes and other mammalian scavengers.
The bone sample consisted of 36 pig (Sus scrofa) femora, obtained commercially. Largely defleshed elements were used to prevent maggot colonization, which could have reduced interest by vertebrate scavengers. To prevent access by the largest scavengers potentially present in this environment, the bones were contained in wire cages (60 cm long, by 43 cm wide, by 50 cm high). The maximum size of gaps in the wire mesh are 10 cm by 14 cm on one side only, and the rest of the openings were smaller than this. Doors to the cages were wired shut. The cages were placed on the ground, with the sample bone wired to the bottom away from the side that has the largest mesh openings, to limit access by larger scavengers attempting to reach in. The cages were staked and wired in place to prevent movement where needed. These test sites were monitored using Bushnell outdoor trail cameras. The cameras are motion-sensitive, use infrared for night vision, and have a picture resolution of 640 X 480 pixels. These were strapped to posts or trees within 2.0 m of each test cage. This distance allowed a full field of view of the cage and its immediate area and was close enough to capture activity if a visiting species interacted with the bone. Where available, some cages had two cameras, with one closer to capture images of smaller species.
Four locations (cages 1-4) were used in mixed forest (cages 1, 3, and 4) and an open, marshy meadow (cage 2) (Figure 1). These were spaced a minimum of 60 m apart to sample multiple territories of individual scavengers and placed within areas of moderate to dense plant cover to prevent disturbance by humans. The bones were placed initially on 8 June 2016, and the images were downloaded every week until 7 October 2016. Each bone showing signs of gnawing or other damage was replaced by a fresh bone, and ungnawed bones also were replaced every two weeks to supply continuous fresh remains for scavenging.
All collected bones were stored in a commercial freezer until field data collection was complete. These were then thawed and examined for any signs of gnawing or other surface marks. The elements displaying surface alteration were dried in a fume hood for multiple weeks, then placed within a dermestid colony maintained by the Boston University School of Medicine Forensic Anthropology Program. Further degreasing was achieved using submersion in an acetone solution. Data were recorded on the location and frequency of gnawing damage type: pits (small, shallow depressions), punctures (small areas penetrating the cortex), scores (long, shallow depressions), furrows (long areas penetrating the cortex), crenellated (irregular) margins, and edge polish (repeated wear of exposed cortical margins) (Pobiner 2008; Pokines 2014). Instances of gnawing damage were correlated with the species observed damaging the bone from the camera images. Each species recorded interacting with the sample bones was identified using standard field guides for mammals and birds inhabiting the Massachusetts/New England region (DeGraaf & Yamasaki 2001; Peterson & Meservey 2004; Veit & Peterson 1993). The data recorded included taxon, number of visits, time of visits, interaction with the bone, and alteration caused to the bones.
Small Scavenger Guild
The taxa recorded by the cameras are presented in Table 1. The total number of observations recorded is 618, with most visits recorded by multiple images of the same individual over a brief span. Multiple taxa showing up in the camera images were never recorded coming into contact with the bone, and these visits totaled only 4.7% of the total number of observations. These include some species of Passerine that might simply have been perching on or adjacent to the cages and represent incidental faunal traffic through the area. A great blue heron (Ardea herodias) also was recorded twice in daytime sightings walking within a camera field. Amphibians observed consisted only of a probable green frog (Lithobates cf. clamitans), and reptiles consisted of snapping turtle (Chelydra serpentina) and garter snake (Thamnophis sirtalis). None of these taxa were noted in contact with the bone and likely were just transiting the area. The snapping turtle was noted passing a cage at night and returning from that same direction the next morning on one day in June, so this individual may have been a female depositing her eggs at night (Congdon et al. 2008) and not attracted to the bone.
Images of multiple mammal species also were captured without any direct contact with the bones recorded during those visits. Groundhog (Marmota monax), white-tailed deer, and gray fox were recorded on multiple visits, and a probable coyote was recorded once. The groundhogs and white-tailed deer showed no interest in the bones and likely were incidental faunal traffic, but the gray fox and probable coyote could have been attracted to the scent of the pig remains. Coyotes likely could not have reached the bones, since the available gaps in the cage bars were relatively small and the bones were not placed close to the margins; these methods were chosen specifically to prevent large scavengers from dominating the scavenging activity. The unknown small mammal sightings were in the size class of rodents.
Multiple bird species were detected interacting with the bone, and the total of all visits from these species (with or without contact) was 18.0% of the total observations. Turkey vultures (Figure 2) were recorded unambiguously feeding on the bones in one location multiple times, as were American crow (Figure 3) and downy woodpecker (Picoides pubescens) (Figure 4). Smaller bird species that were noted contacting the bones consisted of song sparrow (Melospiza melodia), probable dark-eyed junco (cf. Junco hyemalis), probable eastern phoebe (cf. Sayornis phoebe), robin adults and juveniles (Turdus migratorius), house wrens (Troglodytes aedon) (Figure 5), and multiple other unidentified Passerines. Of the identified species, song sparrows were recorded the most times overall (4.5%), followed by house wren (3.5%), although the majority of visits did not include bone contact.
Visits by mammalian taxa that were sometimes observed to contact the pig bones represented the majority of activity recorded (77.3%). These interactions were dominated by a small rodent species, identified as likely deer mouse (Peromyscus maniculatus) or white-footed mouse (P. leucopus). These species are very difficult to tell apart from images alone (Lindquist et al. 2003; Rich et al. 1996), and this was made more difficult by their nocturnal habits (Figure 6). Trapping of specimens and/or examination of owl prey derived from osseous remains in regurgitated pellets in this area may allow identification between these two species. These small rodents accounted for 240 of the 618 total observations (38.8%), with the majority of these (n = 126) including pig bone contact and some clear instances of feeding on external soft tissue. Eastern chipmunk (Tamias striatus) was the next most active mammalian scavenger, with 14.6% of total observations. This species also was noted in multiple instances to be feeding unambiguously on external soft tissue (Figure 7). Other frequent visitors were striped skunk (Mephitis mephitis; 3.4%), long-tailed weasel or stoat (Mustela frenata or M. erminea; 2.6%) (Figure 8), raccoon (4.0%) (Figure 9),Virginia opossum (5.0%) (Figure 10), and eastern gray squirrel (Sciurus carolinensis; 5.0%) (Figure 11). One visit by raccoon consisted of three individuals, which likely represented an adult female and her near-adult offspring foraging together prior to the latter dispersing to their own home territories (Lotze & Anderson 1979). Less frequent visitors were domesticated cat (0.6%), fisher (1.5%) (Figure 12), and northern short-tailed shrew (Blarina brevicauda; 1.0%). The visits by domesticated cat appear to have been all by the same individual and involved two pig bone locations. The taxa identifiable only as unknown medium mammals were in the size class of opossum, raccoon, fisher, or domesticated cat and could have been additional visits by any of these species.
Time of Day
Clear divisions were detected in the timing of visits by multiple taxa, in keeping with their known primary activity patterns (DeGraaf & Yamasaki 2001). Among taxa that contacted the bones, the birds and the domesticated cat were active diurnally. The only exceptions to all bird activity being diurnal were two visits by song sparrow and two visits by unknown Passerines during the crepuscular (twilight) hours. The domesticated cat also may have been restricted to indoors at night. Two rodent species (eastern gray squirrel and chipmunk) were active primarily diurnally, with the exceptions being crepuscular. The fisher was active both night and day, with the daytime visits recorded in full sun. The northern short-tailed shrew and long-tailed weasel/stoat activity was primarily nocturnal, and the daytime visits by these taxa were all crepuscular. The visits by striped skunk, raccoon, Virginia opossum, and deer mouse/white-footed mouse were all nocturnal.
Only the fisher (Figure 12) was observed to gnaw directly upon bone, and two bones bearing distinct traces of carnivore gnawing were recorded. The individual fishers or single fisher that gnawed bones in succession spent several minutes gnawing each and were recorded over multiple images. No other taxa were noted gnawing directly on any of the bones, and no other bones showed signs of surface alteration other than to adhering soft tissue. Rodents do have the potential to gnaw into epiphyses/metaphyses (Haglund 1992, 1997; Pokines 2014; Pokines et al. 2017), so this was examined for in the present study.
Gnawing damage by the fisher was confined to the femoral epiphyses and metaphyses in both cases. Bone #4/6 has no marks on the distal end or shaft (Figure 13). Most of the head was removed, with gnawing extending to the surrounding metaphysis. Part of the greater trochanter also was removed. The margins of the missing bone are irregular/crenellated and generally lack isolated tooth marks along the margins. A small tooth puncture and some furrowing are present on the head (Figures 14 and 15). Bone #1/6 also has no marks on the shaft (Figure 16). Much of the distal condyles was removed, especially the posterior aspect, with irregular/crenellated bone margins created (Figure 17). A small area of gnawing continues onto the posterior right metaphysis. Three tooth punctures are present in the cortical bone of the articular surface of the medial condyle, adjacent to the gnawed margin, and some furrowing is present in the exposed cortical bone. Most of the head and greater trochanter of bone #1/6 was removed, with gnawing damage extending to the surrounding metaphysis (Figure 18). The margins of the missing bone in this area are irregular/crenellated and do not contain discrete tooth marks adjacent. Some furrowing is also present on the head and greater trochanter.
Ricketts (2013), in previous taphonomic research at the ORF using whole juvenile pig carcasses, also recorded scavenging by turkey vultures, Virginia opossum, and fisher. That research further recorded scavenging by black vultures, red-tailed hawk (Buteo jamaicensis), coyote, red fox, and snapping turtle. Junod (2013), also conducting research at the ORF, noted a single scavenging attempt on an isolated, defleshed white-tailed deer bone by a great horned owl (Bubo virginianus) and one instance of bone gnawing by a coyote. Junod (2013) also captured multiple images of white-tailed deer apparently smelling the bones and displaying some interest in them, but no direct contact was recorded. White-tailed deer are known to practice osteophagia (Hutson et al. 2013), including on human bone (Wescott et al. 2016), so bone gnawing by this species is possible but likely rare in this environment. The total number of possible scavenging species compiled through research at the ORF therefore totals 24 (11 bird, 12 mammalian, and 1 reptile), and additional research is likely to increase this total. Potential mammalian scavengers known from Massachusetts and the surrounding region (DeGraaf & Yamasaki 2001; MA EEA 2017b; Sorg et al. 2012) that were not recorded during the present or previous research in this location include red squirrel (Tamiasciurus hudsonicus), northern flying squirrel (Glaucomys sabrinus), southern flying squirrel (Glaucomys volans), bobcat, red fox, black bear, North American river otter (Lontra canadensis), and American mink (Neovison vison) and the introduced species house mouse (Mus musculus), brown/Norway rat (Rattus norvegicus), and domesticated dog. Among these species, only red fox and domesticated dog have been observed directly at the ORF previously. While it is likely that house mouse and brown/Norway rat are present in the area, most of the larger species (black bear, North American river otter, bobcat, and American mink) are likely not currently present in the immediate study area. This situation may change in the future for some of these potential scavengers, as for example with black bears (Lariviere 2001), whose population is on the rise in Massachusetts as they spread from the western portion of the state (MA EEA 2017a). The spread of this larger species also may affect the scavenging opportunities for smaller scavengers, as will the reestablishment of prey species in Massachusetts, including moose (Alces alces) (Foster et al. 2002). Bobcat (Rippley et al. 2012) has been shown elsewhere to be a potential scavenger of human remains, so an increased presence in Massachusetts of this species may affect outdoor forensic scenes and the behavior of other scavengers.
Some rodent species will gnaw wet bone in order to consume the contained soft tissues (Pokines et al. 2017), but no gnawing upon the bones by any of the rodent species was noted during the present research. While rodents often gnaw dry bone for multiple reasons that include wearing down their ever-growing incisors (Haglund 1992, 1997; Klippel & Synstelien 2007; Pokines 2015b; Pokines et al. 2016, 2017; Synstelien 2015), only wet bones were used in the present research. Future research will include direct comparison of wet- versus dry-bone gnawing by rodents in this location.
Gnawing damage in the present research is limited to two instances that may have been caused by the same individual fisher, so no broader comparisons can be drawn. It is noteworthy, however, that only the epiphyses and metaphyses were affected, as it is unlikely that a species this small could effectively attack bones this size in any other manner. It appears that small areas were removed progressively in order to consume the contained grease in the cortical bone. Unlike with larger scavengers, large amounts of separate tooth marks adjacent to the irregular gnawed margins were not created, possibly because the gnawing species in this case cannot open its mouth widely enough to encompass more than a small projecting area of bone at once. The gnawing marks left behind also may have been affected by the bones being secured at midshaft, limiting the distance and angles to which they could be manipulated. Further research with a larger sample may help establish a gnawing profile for fishers and other small scavengers, although much overlap in the size of tooth marks and gnawing behaviors among carnivoran scavenger species has been noted (Delaney-Rivera et al. 2009; Van Valkenburgh 1989).
It is important to note that the present research deliberately excluded the largest scavengers in this environment in order to focus upon the smaller species, so the overall taphonomic effects to individual bones where all potential scavengers are permitted access may be different. It is hoped that the results from this pilot study can shape future research projects at this forensic facility, including direct comparisons of dry-bone versus wet-bone gnawing by rodents and the timing these behaviors manifest during the postmortem interval (Klippel & Synstelien 2007; Pokines 2014, 2015b; Pokines et al. 2016, 2017). The present research also noted bone gnawing in only two instances and by the same species, fisher. The greatest overall effect that small scavengers have upon sets of terrestrially deposited skeletal remains, however, could be through dispersal of elements away from their initial point of deposition (Pokines 2014), especially given that the present study noted little alteration of the bones themselves. Only long bones were used, so it is possible that bones with smaller margins such as vertebrae and ribs may be more suitable for gnawing by smaller species. Multiple rodent species are known to disperse bone (Hockett 1989; Haglund et al. 1989; Hoffman & Hays 1987; Klippel & Synstelien 2007; Synstelien 2015), so their actions may greatly increase the search areas necessary to process outdoor forensic scenes. Future research into this topic may aid in the planning and implementation of forensic scene processing (Pokines 2015a) and may include human remains, clothed and unclothed.
Due to potential repeat baiting effects whereby the animals might become accustomed to returning to the same food source repeatedly (Stringham 1986; Synstelien 2015), the present research cannot quantitatively assess the relative frequency and overall impact of the small vertebrate scavengers in this area under more natural conditions. Similarly, the territorial behavior (Bekoff 1977; Lariviere & Pasitschniak-Arts 1996; McManus 1974; Way 2007) of individual animals also may affect their frequency of discovering food sources introduced into their territories. The present research combined with previous taphonomic research conducted at the Holliston ORF (Junod 2013; Ricketts 2013) does indicate that a broad spectrum of bird and mammal scavengers are active in this environment, despite the proximity of human habitations to a wooded area within a town with an estimated 14,525 inhabitants distributed over 18.64 square miles, or 726.6 persons/square mile (U.S. Census Bureau 2017). The most frequent scavengers were noted to be mice (Peromyscus sp.) and eastern chipmunks, although it is unlikely that they are responsible for the majority of soft-tissue loss from the experimental bones given the small body mass of these taxa. It is also clear that diurnal and nocturnal scavenging activity is typical, and a single bone may be used as a resource by multiple individual animals. The effects of these species is likely widespread and pervasive for all sets of human remains deposited outdoors over a sufficient duration, and their activity can continue long after most of the soft tissue was consumed by blowfly maggot masses or larger scavengers. After most soft tissue, including grease content, has decomposed, the skeletal elements will continue to attract activity from rodents gnawing upon the dry bone (Pokines 2015b, 2016; Pokines et al. 2016, 2017).
Due to the inherent skittishness of the species examined, their small sizes, and the large amount of nocturnal scavenging, the authors recommend the use of wildlife cameras to record their scavenging activity. The system used in the present research also allowed for the identification of small diurnal bird species. In addition, the difficulty of detection of many of the taxa examined here indicates their potential to alter the results of decomposition studies where the research protocol was designed to exclude large scavengers but was insufficient to exclude small scavengers (Synstelien 2015). Loss of soft tissue through scavenging may affect the results of decomposition sequences used to estimate the postmortem interval (Suckling et al. 2016).
The taxa reported here are among the probable scavengers in this environment of human remains deposited in non-urban, outdoor environments and therefore of considerable forensic significance in terms of soft-tissue scavenging and, to a more limited extent, bone gnawing. Continued research in this location may indicate an even broader small scavenger guild, but it is likely that most of the current common mammalian taxa for this area have been identified, given the known species of Massachusetts (MA EEA 2017b). As noted, the potential for territorial re-expansion may alter this species community composition in the future (MA EEA 2017a). Testing in different environments, including more open territory, different forest types, and the coastal region, may expand significantly the breadth of the bird component of the small scavenger guild (Peterson & Meservey 2004). Additional research also should focus on the long-term and seasonal effects of these species upon terrestrial surface remains, individual species' taphonomic effects (including gnawing upon fresh bone), and dispersal of remains from their points of initial deposition (Kjorlien 2004; Kjorlien et al. 2009). Since many of these species or their close relatives have ranges that extend far beyond Massachusetts, their taphonomic effects likely extend across wider regions in North America and require additional research in other locations.
The authors thank the Boston University School of Medicine, Forensic Anthropology Program for the use of the Outdoor Research Facility in Holliston, Massachusetts. The authors also thank Dr. Eric Bartelink and the two anonymous peer reviewers for their valuable comments on earlier versions of this manuscript.
Bekoff M. Canis latrans. Mammalian Species. 1977;79:1-9.
Congdon JD, Greene JL, Brooks RJ. Reproductive and nesting ecology of female snapping turtles. In: Steyermark AC, Finkler MS, Brooks RJ, eds. Biology of the Snapping Turtle (Chelydra serpentina). Baltimore: Johns Hopkins University Press; 2008:124-134.
Cortes-Avizanda A, Jovani R, Carrette M, Donazar JA. Resource unpredictability promotes species diversity and coexistence in an avian scavenger guild: A field experiment. Ecology 2012;93(12):2570-2579.
DeGraaf RM, Yamasaki M. New England Wildlife: Habitat, Natural History, and Distribution. Hanover, NH: University Press of New England; 2001.
DeGraaf RM, Yamasaki M, Leak WB, Lester AM. Technical Guide to Forest Wildlife Habitat Management in New England. Burlington: University of Vermont Press; 2006.
Delaney-Rivera C, Plummer TW, Hodgson JA, Forrest F, Hertel F, Oliver JS. Pits and pitfalls: Taxonomic variability and patterning in tooth mark dimensions. Journal of Archaeological Science 2009;36(11):2597-2608.
Foster DR, Motzkin G, Bernardos D, Cardoza J. Wildlife dynamics in the changing New England landscape. Journal of Biogeography 2002;29:337-1357.
Haglund WD. Contribution of rodents to postmortem artifacts of bone and soft tissue. Journal of Forensic Sciences 1992;37(6): 1459-1465.
Haglund WD. Rodents and human remains. In: Haglund WD, Sorg MH, eds. Forensic Taphonomy: The Postmortem Fate of Human Remains. Boca Raton, FL: CRC Press; 1997:405-414.
Haglund WD, Reay DT, Swindler DR. Tooth mark artifacts and survival of bones in animal scavenged human skeletons. Journal of Forensic Sciences 1988;33(4):985-997.
Haglund WD, Reay DT, Swindler DR. Canid scavenging/disarticulation sequence of human remains in the Pacific Northwest. Journal of Forensic Sciences 1989;34(3):587-606.
Haynes G. A guide to differentiating mammalian carnivore taxa responsible for gnaw damage to herbivore limb bones. Paleobiology 1983;9(2):164-172.
Hockett BS. The concept of "carrying range": A method for determining the role played by woodrats in contributing bones to archaeological sites. Nevada Archaeologist 1989;7(1):28-35.
Hoffman R, Hays C. The eastern wood rat (Neotoma floridana) as a taphonomic factor in archaeological sites. Journal of Archaeological Science 1987;14:325-337.
Hutson JM, Burke CC, Haynes G. Osteophagia and bone modifications by giraffe and other large ungulates. Journal of Archaeological Science 2013;40;4139-4149.
Jeong Y, Meadows Jantz L, Smith J. Investigation into seasonal scavenging patterns of raccoons on human decomposition. Journal of Forensic Sciences 2016;61(2):467-471.
Junod CA. Subaerial Bone Weathering and Other Taphonomic Changes in a Temperate Climate [master's thesis]. Boston: Boston University School of Medicine; 2013.
Kjorlien YP. Patterns in the Scattering of Remains due to Scavenger Activity [master's thesis]. Edmonton, Canada: University of Alberta; 2004.
Kjorlien YP, Beattie OB, Peterson AE. Scavenging activity can produce predictable patterns in surface skeletal remains scattering: Observations and comments from two experiments. Forensic Science International 2009;188:103-106.
Klippel WE, Synstelien JA. Rodents as taphonomic agents: Bone gnawing by brown rats and gray squirrels. Journal of Forensic Sciences 2007;52(4):765-773.
Komar D, Beattie O. Identifying bird scavenging in fleshed and dry remains. Canadian Society of Forensic Science Journal 1998;31(3):177-188.
Lariviere S. Ursus americanus. Mammalian Species. 2001;647:1-11.
Lariviere S, Pasitschniak-Arts M. Vulpes vulpes. Mammalian Species. 1996;537:1-11.
Lindquist ES, Aquadro CF, McClearn D, McGowan KJ. Field identification of the mice Peromyscus leucopus noveboracensis and P. maniculatus gracilis in central New York. Canadian Field Naturalist 2003;117:184-189.
Lotze J-H, Anderson S. Procyon lotor. Mammalian Species 1979; 119:1-8.
MA EEA [Massachusetts Energy and Environmental Affairs]. Black bears in MA. http://www.mass.gov/eea/agencies/dfg/dfw/fish-wildlife-plants/mammals/black-bear-mass.html. Accessed February 19, 2017a.
MA EEA [Massachusetts Energy and Environmental Affairs]. State mammal list. http://www.mass.gov/eea/agencies/dfg/dfw/fish-wildlife-plants/state-mammal-list.html. Accessed February 19, 2017b.
McManus JJ. Didelphis virginiana. Mammalian Species 1974;40: 1-6.
Morton RJ, Lord WD. Taphonomy of child-sized remains: A study of scattering and scavenging in Virginia, USA. Journal of Forensic Sciences 2006;51(3):475-479.
Moss KE. The Effects of Avian and Terrestrial Scavenger Activity on Human Remains and Decomposition in Southeast Texas During an 18 Month Study [master's thesis]. Houston: University of Houston; 2012.
Murmann DC, Brumit PC, Schrader BA, Senn DR. A comparison of animal jaws and bite mark patterns. Journal of Forensic Sciences 2006;51(4):846-860.
O'Brien RC, Forbes SL, Meyer J, Dadour I. Forensically significant scavenging guilds in the southwest of Western Australia. Forensic Science International 2010;198:85-91.
Olson ZH, Beasley JC, Rhodes OE, Jr. Carcass type affects local scavenger guilds more than habitat connectivity. PLoS One 2016;11(2):e0147798. DOI: 10.1371/journal.pone. 0147798.
Peterson WR, Meservey WR, eds. Massachusetts Breeding Bird Atlas. Amherst: Massachusetts Audubon Society; 2004.
Pobiner B. Paleoecological information in predator tooth marks. Journal of Taphonomy 2008;6(3-4):373-397.
Pokines JT. Faunal dispersal, reconcentration, and gnawing damage to bone in terrestrial environments. In: Pokines JT, Symes SA, eds. Manual of Forensic Taphonomy. Boca Raton, FL: CRC Press; 2014:201-248.
Pokines JT. A procedure for processing outdoor surface forensic scenes yielding skeletal remains among leaf litter. Journal of Forensic Identification 2015a;65(2):161-172.
Pokines JT. Taphonomic alterations by the rodent species woodland vole (Microtus pinetorum) upon human skeletal remains. Forensic Science International 2015b;257:e16-e19.
Pokines JT. Taphonomic alterations to terrestrial surface-deposited human osseous remains in a New England, U.S.A. environment. Journal of Forensic Identification 2016;66(1):59-78.
Pokines JT, Baker SE. Avian taphonomy. In: Pokines JT, Symes SA, eds. Manual of Forensic Taphonomy. Boca Raton, FL: CRC Press; 2014:427-446.
Pokines JT, Santana SA, Hellar JD, Bian P, Downs A, Wells N, Price MD. The taphonomic effects of eastern gray squirrel (Sciurus carolinensis) gnawing upon bone. Journal of Forensic Identification 2016;66(4):349-375.
Pokines JT, Sussman R, Gough M, Ralston C, McLeod E, Brun K, Kearns A, Moore TL. Taphonomic analysis of Rodentia and Lagomorpha bone gnawing based upon incisor size. Journal of Forensic Sciences 2017;62(1):50-66.
Reeves NM. Taphonomic effects of vulture scavenging. Journal of Forensic Sciences 2009;54(3):523-528.
Ricketts DR. Scavenging Effects and Scattering Patterns on Porcine Carcasses in Eastern Massachusetts [master's thesis]. Boston: Boston University School of Medicine; 2013.
Rich SM, Kilpatrick CW, Shippee JL, Crowell KL. Morphological differentiation and identification of Peromyscus leucopus and P. maniculatus in northeastern North America. Journal of Mammalogy 1996;77(4):985-991.
Rippley A, Larison NC, Moss KE, Kelly JD, Bytheway JA. Scavenging behavior of Lynx rufus on human remains during the winter months of southeast Texas. Journal of Forensic Sciences 2012;57(3):699-705.
Root RB. The niche exploitation pattern of the blue-gray gnat-catcher. Ecological Monographs 1967;37:317-350.
Simberloff D, Dayan T. The guild concept and the structure of ecological communities. Annual Review of Ecology, Evolution and Systematics 1991;22:115-143.
Smith AC. The effects of sharp-force thoracic trauma on the rate and pattern of decomposition. Journal of Forensic Sciences 2014;59(2):319-326.
Sorg MH, Haglund WD, Wren JA, Collar A. Taphonomic impacts of small and medium-sized scavengers in northern New England. Proceedings of the 64th Annual Meeting of the American Academy of Forensic Sciences, February 20-25, 2012; Atlanta, GA.
Spradley MK, Hamilton MD, Giordano A. Spatial patterning of vulture scavenged human remains. Forensic Science International 2012;219:57-63.
Stolen ED. Turkey vultures carrying carrion. Florida Field Naturalist 2003;31:2.
Stringham SF. Effects of climate, dump closure, and other factors on Yellowstone grizzly bear litter size. International Conference on Bear Research and Management 1986;6:33-39.
Suckling JK, Spradley MK, Godde K. A longitudinal study on human outdoor decomposition in central Texas. Journal of Forensic Sciences 2016;61(2):19-25.
Synstelien JA. Studies in Taphonomy: Bone and Soft Tissue Modifications by Postmortem Scavengers [PhD dissertation]. Knoxville: University of Tennessee; 2015.
U.S. Census Bureau. http://www.census.gov/en.html. Accessed February 19, 2017.
Van Valkenburgh BV. Carnivore dental adaptations and diet: A study of trophic diversity within guilds. In: Gittleman JL, ed. Carnivore Behavior, Ecology, and Evolution. Vol 1. London: Chapman & Hall; 1989:410-436.
Veit RR, Peterson WR. Birds of Massachusetts. Lincoln, MA: Massachusetts Audubon Society; 1993.
Wallace MP, Temple SA. Competitive interactions within and between species in a guild of avian scavengers. The Auk 1987;104(2):290-295.
Way JG. Suburban Howls: Tracking the Eastern Coyote in Eastern Massachusetts. Indianapolis, IN: Dog Ear Publications; 2007.
Wescott DJ, Meckel LA, McDaneld CP, Hamilton MD, Mavroudas S, Spradley K. White-tailed deer as a taphonomic agent: Photographic documentation of white-tailed deer gnawing on human bone. Proceedings of the 68th Annual Meeting of the American Academy of Forensic Sciences, February 22-27, 2016; Las Vegas, NV.
Young A, Marquez-Grant N, Stillman R, Smith MJ, Korstjens AH. An investigation of red fox (Vulpes vulpes) and Eurasian badger (Meles meles) scavenging, scattering, and removal of deer remains: Forensic implications and applications. Journal of Forensic Sciences 2015;60(S1):S39-S55.
Young A, Stillman R, Smith MJ, Korstjens AH. Applying knowledge of species-typical scavenging behavior to the search and recovery of mammalian skeletal remains. Journal of Forensic Sciences 2016;60(2):458-456.
Young A, Stillman R, Smith MJ, Korstjens AH. An experimental study of vertebrate scavenging behavior in a northwest European woodland context. Journal of Forensic Sciences 2014;59(5):1333-1342.
James Pokines (a,b*) * Corey Pollock (a)
(a) Department of Anatomy and Neurobiology, Boston University School of Medicine, USA
(b) Office of the Chief Medical Examiner, Boston, MA, USA
(*) Correspondence to: James Pokines, Department of Anatomy and Neurobiology, Boston University School of Medicine, 72 E. Concord Street, Boston, MA 02118, USA
Received 19 February 2017; Revised 22 March 2017; Accepted 15 April 2017
TABLE 1--Results of scavenging monitoring, 8 June to 7 October 2016, ORF, Holliston, Massachusetts. Day with Common Name Taxon Contact Taxa Not Detected Contacting Bone Birds Great blue heron Ardea herodias 0 Gray catbird Dumetella carolinensis 0 Common yellowthroat Geothlypis trichas 0 Black-capped chickadee Poecile atricapillus 0 Common grackle cf. Quiscalus quiscula 0 Common starling cf. Sturnus vulgaris 0 Mammals Coyote cf. Canis latrans 0 Groundhog Marmota monax 0 White-tailed deer Odocoileus virginianus 0 Gray fox Urocyon cinereoargenteus 0 Unknown mammal 0 Unknown small mammal 0 Amphibians/Reptiles Green frog Lithobates cf. clamitans 0 Snapping turtle Chelydra serpentina 0 Garter snake Thamnophis sirtalis 0 Unknown vertebrate 0 Taxa Detected Contacting Bone Birds Song sparrow Melospiza melodia 0 American crow Corvus brachyrhynchos 1 Dark-eyed junco cf. Junco hyemalis 1 Eastern phoebe cf. Sayornis phoebe 1 Robin Turdus migratorius 2 Turkey vulture Cathartes aura 4 House wren Troglodytes aedon 4 Downy woodpecker Picoides pubescens 6 Unknown Passerine 2 Mammals Virginia opossum Didelphis virginiana 0 Northern short-tailed shrew Blarina brevicauda 2 Striped skunk Mephitis mephitis 0 Long-tailed weasel/stoat Mustela frenata/erminea 0 Fisher Martes pennanti 3 House cat Felis catus 1 Raccoon Procyon lotor 0 Deer mouse/white-footed mouse cf. Peromyscus maniculatus/leucopus 0 Eastern gray squirrel Sciurus carolinensis 1 Eastern chipmunk Tamias striatus 55 Unknown medium mammal 0 Grand Totals 83 Night with Day no Night no Common Name Contact Contact Contact Taxa Not Detected Contacting Bone Birds Great blue heron 0 2 0 Gray catbird 0 2 0 Common yellowthroat 0 1 0 Black-capped chickadee 0 1 0 Common grackle 0 1 0 Common starling 0 2 0 Mammals Coyote 0 0 1 Groundhog 0 4 0 White-tailed deer 0 4 1 Gray fox 0 1 1 0 0 1 0 1 1 Amphibians/Reptiles Green frog 0 1 0 Snapping turtle 0 1 1 Garter snake 0 1 0 Unknown vertebrate 0 0 1 Taxa Detected Contacting Bone Birds Song sparrow 1 26 1 American crow 0 2 0 Dark-eyed junco 0 0 0 Eastern phoebe 0 2 0 Robin 0 9 0 Turkey vulture 0 0 0 House wren 0 18 0 Downy woodpecker 0 2 0 Unknown Passerine 1 27 1 Mammals Virginia opossum 16 0 15 Northern short-tailed shrew 2 0 2 Striped skunk 6 0 15 Long-tailed weasel/stoat 10 1 5 Fisher 4 1 1 House cat 0 3 0 Raccoon 10 0 15 Deer mouse/white-footed mouse 126 0 114 Eastern gray squirrel 0 29 1 Eastern chipmunk 0 30 5 Unknown medium mammal 1 0 4 177 172 186 Total % Total Common Name Visits Visits Taxa Not Detected Contacting Bone Birds Great blue heron 2 0.3 Gray catbird 2 0.3 Common yellowthroat 1 0.2 Black-capped chickadee 1 0.2 Common grackle 1 0.2 Common starling 2 0.3 Mammals Coyote 1 0.2 Groundhog 4 0.6 White-tailed deer 5 0.8 Gray fox 2 0.3 1 0.2 2 0.3 Amphibians/Reptiles Green frog 1 0.2 Snapping turtle 2 0.3 Garter snake 1 0.2 Unknown vertebrate 1 0.2 Total 4.7 Taxa Detected Contacting Bone Birds Song sparrow 28 4.5 American crow 3 0.5 Dark-eyed junco 1 0.2 Eastern phoebe 3 0.5 Robin 11 1.8 Turkey vulture 4 0.6 House wren 22 3.6 Downy woodpecker 8 1.3 Unknown Passerine 31 5.0 Total 18.0 Mammals Virginia opossum 31 5.0 Northern short-tailed shrew 6 1.0 Striped skunk 21 3.4 Long-tailed weasel/stoat 16 2.6 Fisher 9 1.5 House cat 4 0.6 Raccoon 25 4.0 Deer mouse/white-footed mouse 240 38.8 Eastern gray squirrel 31 5.0 Eastern chipmunk 90 14.6 Unknown medium mammal 5 0.8 Total 77.3 618
|Printer friendly Cite/link Email Feedback|
|Title Annotation:||RESEARCH ARTICLE|
|Author:||Pokines, James; Pollock, Corey|
|Date:||Jan 1, 2018|
|Previous Article:||Forensic Fractography of Bone A New Approach to Skeletal Trauma Analysis.|
|Next Article:||Using an Alternate Light Source (ALS) to Search for Surface Deposited Skeletal Remains.|